This article provides insufficient context for those unfamiliar with the subject.(January 2023) |
Candidate phyla radiation | |
---|---|
Drawing of a CPR bacterium from a "GWB1" sample. | |
Scientific classification | |
Domain: | Bacteria |
(unranked): | Bacteria candidate phyla |
Infrakingdom: | Candidate phyla radiation |
The candidate phyla radiation (also referred to as CPR group) is a large evolutionary radiation of bacterial lineages whose members are mostly uncultivated and only known from metagenomics and single cell sequencing. They have been described as nanobacteria (not to be confused with non-living nanoparticles of the same name) or ultra-small bacteria due to their reduced size (nanometric) compared to other bacteria.
Originally (circa 2016), it has been suggested that CPR represents over 15% of all bacterial diversity and may consist of more than 70 different phyla. [1] However, the Genome Taxonomy Database (2018) based on relative evolutionary divergence found that CPR represents a single phylum, [2] with earlier figures inflated by the rapid evolution of ribosomal proteins. [3] CPR lineages are generally characterized as having small genomes and lacking several biosynthetic pathways and ribosomal proteins. This has led to the speculation that they are likely obligate symbionts. [4] [5]
Earlier work proposed a superphylum called Patescibacteria which encompassed several phyla later attributed to the CPR group. [6] Therefore, Patescibacteria and CPR are often used as synonyms. [7] The former name is not necessarily obsolete: for example, the GTDB uses this name because they consider the CPR group a phylum. [2]
Although there are a few exceptions, members of the candidate phyla radiation generally lack several biosynthetic pathways for several amino acids and nucleotides. To date, there has been no genomic evidence that indicates that they are capable of producing the lipids essential for cell envelope formation. [5] Additionally, they tend to lack complete TCA cycles and electron transport chain complexes, including ATP synthase. This lack of several important pathways found in most free-living prokaryotes indicates that the candidate phyla radiation is composed of obligate fermentative symbionts. [8]
Furthermore, CPR members have unique ribosomal features. While the members of CPR are generally uncultivable, and therefore missed in culture-dependent methods, they are also often missed in culture-independent studies that rely on 16S rRNA sequences. Their rRNA genes appear to encode proteins and have self-splicing introns, features that are rarely seen in bacteria, although they have previously been reported. [9] Owing to these introns, members of CPR are not detected in 16S-dependent methods. Additionally, all CPR members are missing the L30 ribosomal protein, a trait that is often seen in symbionts. [8]
Many of its characteristics are similar or analogous to those of ultra-small archaea (DPANN). [5]
The Candidate phyla radiation was found to be the most basal-branching lineage in bacteria according to some early phylogenetic analyses of this group based on ribosomal proteins and protein family occurrence profiles. These studies found the following phylogeny between phyla and superphyla. The superphyla are shown in bold. [5] [4]
Bacteria |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
However, several recent studies have suggested that the CPR belongs to Terrabacteria and is more closely related to Chloroflexota. [11] [12] [13] The evolutionary relationships that are typically supported by these studies are as follows.
Bacteria |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||
Because many CPR members are uncultivable, they cannot be formally put into the bacterial taxonomy, but a number of provisional, or Candidatus , names have been generally agreed on. [6] [14] As of 2017, two superphyla are generally recognized under CPR, Parcubacteria and Microgenomates. [1] The Phyla under CPR include:
Phylogeny of Patescibacteria [15] [16] [17] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
|
Phylogeny of Microgenomatia [15] [16] [17] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
|
Phylogeny of Gracilibacteria [15] [16] [17] | |||||||||||||||||||||||||||||||||
|
Phylogeny of ABY1 [15] [16] [17] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||
|
Phylogeny of Paceibacteria [15] [16] [17] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
|
The current phylogeny is based on ribosomal proteins (Hug et al., 2016). [4] Other approaches, including protein family existence and 16S ribosomal RNA, produce similar results at lower resolutions. [18] [1]
Neomura is a possible clade composed of the two domains of life of Archaea and Eukaryota. The group was named by Thomas Cavalier-Smith in 2002. Its name means "new walls", reflecting his hypothesis that it evolved from Bacteria, and one of the major changes was the replacement of peptidoglycan cell walls with other glycoproteins. As of August 2017, the neomuran hypothesis is not accepted by most workers; molecular phylogenies suggest that eukaryotes are most closely related to one group of archaeans and evolved from them, rather than forming a clade with all archaeans, and that Archaea and Bacteria are sister groups.
Gracilicutes is a clade in bacterial phylogeny.
The PVC superphylum is a superphylum of bacteria named after its three important members, Planctomycetota, Verrucomicrobiota, and Chlamydiota. Cavalier-Smith postulated that the PVC bacteria probably lost or reduced their peptidoglycan cell wall twice. It has been hypothesised that a member of the PVC clade might have been the host cell in the endosymbiotic event that gave rise to the first proto-eukaryotic cell.
Archaeal Richmond Mine acidophilic nanoorganisms (ARMAN) were first discovered in an extremely acidic mine located in northern California (Richmond Mine at Iron Mountain) by Brett Baker in Jill Banfield's laboratory at the University of California Berkeley. These novel groups of archaea named ARMAN-1, ARMAN-2 (Candidatus Micrarchaeum acidiphilum ARMAN-2), and ARMAN-3 were missed by previous PCR-based surveys of the mine community because the ARMANs have several mismatches with commonly used PCR primers for 16S rRNA genes. Baker et al. detected them in a later study using shotgun sequencing of the community. The three groups were originally thought to represent three unique lineages deeply branched within the Euryarchaeota, a subgroup of the Archaea. However, based on a more complete archaeal genomic tree, they were assigned to a new superphylum named DPANN. The ARMAN groups now comprise deeply divergent phyla named Micrarchaeota and Parvarchaeota. Their 16S rRNA genes differ by as much as 17% between the three groups. Prior to their discovery, all of the Archaea shown to be associated with Iron Mountain belonged to the order Thermoplasmatales (e.g., Ferroplasma acidarmanus).
Terrabacteria is a taxon containing approximately two-thirds of prokaryote species, including those in the gram positive phyla as well as the phyla "Cyanobacteria", Chloroflexota, and Deinococcota.
The FCB group is a superphylum of bacteria named after the main member phyla Fibrobacterota, Chlorobiota, and Bacteroidota. The members are considered to form a clade due to a number of conserved signature indels.
Bacterial phyla constitute the major lineages of the domain Bacteria. While the exact definition of a bacterial phylum is debated, a popular definition is that a bacterial phylum is a monophyletic lineage of bacteria whose 16S rRNA genes share a pairwise sequence identity of ~75% or less with those of the members of other bacterial phyla.
Saccharibacteria, formerly known as TM7, is a major bacterial lineage. It was discovered through 16S rRNA sequencing.
Hydrobacteria is a taxon containing approximately one-third of prokaryote species, mostly gram-negative bacteria and their relatives. It was found to be the closest relative of an even larger group of Bacteria, Terrabacteria, which are mostly gram positive bacteria. The name Hydrobacteria refers to the moist environment inferred for the common ancestor of those species. In contrast, species of Terrabacteria possess adaptations for life on land.
DPANN is a superphylum of Archaea first proposed in 2013. Many members show novel signs of horizontal gene transfer from other domains of life. They are known as nanoarchaea or ultra-small archaea due to their smaller size (nanometric) compared to other archaea.
The Orthokaryotes are a proposed Eukaryote clade consisting of the Jakobea and the Neokaryotes. Together with its sister Discicristata it forms a basal Eukaryote clade. They are characterized by stacked Golgi, orthogonal centrioles, and two opposite posterior ciliary roots.
The neokaryotes are a proposed eukaryote clade consisting of the unikonts and the bikonts as sister of for instance the Jakobea. It arises because the Euglenozoa, Percolozoa, Tsukubea, and Jakobea are seen in this view as more basal eukaryotes. These four groups, are traditionally grouped together in the Discoba. However, the Discoba may well be paraphyletic as the neokaryotes may have emerged in them.
The Scotokaryotes (Cavalier-Smith) is a proposed basal Neokaryote clade as sister of the Diaphoretickes. Basal Scotokaryote groupings are the Metamonads, the Malawimonas and the Podiata. In this phylogeny the Discoba are sometimes seen as paraphyletic and basal Eukaryotes.
Asgard or Asgardarchaeota is a proposed superphylum consisting of a group of archaea that contain eukaryotic signature proteins. It appears that the eukaryotes, the domain that contains the animals, plants, and fungi, emerged within the Asgard, in a branch containing the Heimdallarchaeota. This supports the two-domain system of classification over the three-domain system.
The Microgenomates are a proposed supergroup of bacterial candidate phyla in the Candidate Phyla Radiation.
Gracilibacteria is a bacterial candidate phylum formerly known as GN02, BD1-5, or SN-2. It is part of the Candidate Phyla Radiation and the Patescibacteria group.
Katanobacteria is a bacterial phylum formerly known as WWE3. It has candidate status, meaning there are no cultured representatives, and is a member of the Candidate Phyla Radiation (CPR).
CandidatusWirthbacteria is a proposed bacterial phylum containing only one known sample from the Crystal Geyser aquifer, Ca. Wirthibacter wanneri. This bacterium stands out in a basal position in some trees of life as it is closely related to Candidate phyla radiation but is not considered part of that clade.
Nomurabacteria is a candidate phylum of bacteria belonging to the CPR group so they are ultra-small bacteria. They have been found in a wide variety of environments, mainly in sediments under anaerobic conditions.
There are several models of the branching order of bacterial phyla, one of these is the Genome Taxonomy Database (GTDB).